Alternative mating strategy

An alternative mating strategy is a mating strategy used

by males or females that diers from the prevailing strategy of the sex. The mating strategies of animals are diverse and variable both across and within species. Animal
sexual behavior and mate choice directly aect social
structure and relationships in many dierent mating systems, whether monogamous, polygamous, polyandrous,
or polygynous. Though males and females in a given
population typically employ a predominant reproductive
strategy based on the overarching mating system, there is
still signicant variation in behavior among individuals
of the same sex.[1] Alternative strategies provide animals
of certain phenotypes with a dierent means for obtaining mates.[1] The study of alternative mating strategies is
critical to correctly characterizing the diverse sexual behavior practiced by animals in a population and understanding the strength of selection on these individuals.

for phenotypes and strategies that maximize an animals

chance of obtaining a mate. As a result, certain animals
will successfully use a conventional mating strategy while
others employing this strategy will not obtain mates. Over
time, phenotypic variance both between and within the
sexes will result, with males exhibiting greater diversity
in phenotype.[2] The resulting variance in male tness will
create a niche in which alternative strategies may develop,
such as sneaking to obtain a mate. The alternative behaviors will persist as part of this polymorphism, or variety
of phenotypes, because the average tness of unconventional males will equal the average reproductive success
of conventional males.[2]
Alternative behaviors will be maintained through
frequency-dependent selection because of their equal
tness benets and functional equivalence.[3] Under
frequency-dependent selection, the tness of a given
phenotype is determined by its frequency relative to
other phenotypes within a population. Similarly, negative frequency-dependent selection describes a scenario
in which rarer phenotypes experience greater tness.[4]
Given that the utilization of alternative mating strategies
has been shown to uctuate over time, it has been suggested that frequency or negative frequency-dependent
selection is the mechanism through which alternative
mating strategies are maintained in animal populations.[4]

Strategies and selection

Alternative mating strategies have been observed among

both male and female animals.[2] Most typically, alternative strategies will be adopted in the face of competition
within a sex, especially in species that mate multiply. In
these scenarios, some individuals will adopt very dierent mating strategies to achieve reproductive success.[3]
The result over time will be a variety of strategies and
phenotypes, consisting of both unsuccessful and successful conventional individuals and unconventional individuals who mate through alternative means. Successful
strategies will be maintained through sexual selection.

s"
A"
B"

In many cases, the coexistence of alternative and traditional mating strategies will both maximize the average
tness of the sex in question and be evolutionarily stable
for a population.[2] However, the utilization of alternative mating strategies may oscillate as a result of varying
reproductive conditions, such as the availability of potential mates. Under changing circumstances, the existence
of a variety of strategies allows individuals to choose the
conditional behavior that will currently maximize their
tness.[1]

1.1

Figure 1: This graph describes the tness payos of varying mating strategies in relation to an animals status. At the intermediate
point s, the tness benets of A and B are equal, and either phenotype may be expressed. Individuals of higher status above point
s will receive more tness benets more from exhibiting phenotype A, while those of lower status will achieve higher tness with
phenotype B.

Selection

Conventional and alternative mating behaviors arise A second proposed model for the maintenance of alternathrough the pressures of sexual selection. More specif- tive mating behaviors is status-dependent selection. This
ically, varying levels of reproductive success will select describes a conditional strategy in which the tness of al1

ternative phenotypes depend on the status, or competitive

ability, of an individual.[1] Status includes environmental and genetic factors as well as age and size, and determines the level of tness that may be obtained from a
given phenotype.[1] As shown in Figure 1, the tness benets of a given phenotype vary based on whether an individual is of high or low status. In a case where two phenotypes and strategies are possible, such as mate guarding or sneaking, there will be an intermediate point of
intersection where the tness gained from these alternative behaviors will be equivalent. At this point (s), the
tness gained from these strategies will be equal, and the
particular strategy employed at a given time will depend
on an individuals status.[1] A low status individual below the switch point will obtain higher tness with phenotype B, while an individual of high status above the
switch point will benet from higher tness with phenotype A. Such a model shows how individuals of lesser
status or competitive ability may maximize their tness
by exhibiting an alternative phenotype. In this manner,
these selective forces will maintain the phenotypic diversity observed among animals with respect to mating behavior, though strategies utilized will depend on a variety
of circumstances.

1.2

Strategy

Most of the organisms in question do not have the cognitive capacity to strategize in the human sense of the
word, so what is a strategy? Here, a strategy is an underlying rule for making decisions about a certain behavior. A
strategy provides an organism with a set of tactics that are
adaptive in various circumstances. A tactic is an action
taken to achieve a specic goal.[2] For example, a wolf
encounters a fallen tree and its strategy is dened by two
tactics that may allow the wolf to pass the obstacle: jump
over it or crawl under it. Considering the current environmental conditions, the surroundings, and the size of
the tree, the wolf will decide between the tactics dictated
by its strategy. In the context of a mating system, this
means that individuals in a given population have strategies that allow them to obtain mates in dierent ways to
maximize their reproductive success given their phenotypic, environmental, or social circumstances.
It is important to recognize that organisms within a population may not always have the same strategy, and different strategies may oer individuals either a range of
tactical options or just one tactic. Furthermore, given
strategy may be considered Mendelian, developmental,
conditional, or a combination of the above. A Mendelian
strategy depends on a genetically determined phenotypic dierence, such as body size. This is the case
in marine isopods, described below. Developmentally
driven strategies are associated with phenotypic dierences caused by varying conditions during the course of
development that aect body size or overall adult health.
Individuals may also have a conditional behavior strategy

STRATEGIES AND SELECTION

that depends not on the genetic or developmental impact

on ones life circumstance, but on external factors. These
may include the number of available mates, or the number
of nearby competitors and their employed tactics. Additionally, some mating strategies will be impacted by the
interaction of multiple factors, so these categorizations of
Mendelian, developmental, and conditional are not mutually exclusive. They simply oer ways to think about
alternative mating strategies and their root causes.[2]
In any case, the mating strategies employed by organisms in various situations will ultimately depend on the
strength of selection acting to maintain or eliminate certain reproductive strategies. If sexual selection strongly
favors one mating strategy over a potential alternative,
individuals not conforming to the successful strategy will
fail to reproduce, thus preventing future generations from
inheriting the unsuccessful strategy.[2]

1.3 Evolutionarily stable strategy

The diversity of mating strategies within animal populations may be understood through evolutionary game
theory concepts that assess the costs and benets of reproductive decision-making. The Evolutionarily Stable
Strategy (ESS) concept provides a particularly useful
framework for considering alternative behaviors as they
relate to tness. Given that a strategy describes a set of
pre-programmed rules that specify particular behaviors,
an evolutionarily stable strategy is one that persists in a
population due to its benets to tness.[3] An ESS will be
maintained in a population if it accords higher average
tness than other strategies, or a level of average individual tness equivalent to all other strategies within the
population.[2]
Within an evolutionarily stable strategy, several scenarios
are possible, including pure and mixed strategies. A pure
strategy is one not aected by chance, in which an individual only expresses one strategic behavior.[1] In contrast, a mixed strategy describes a scenario involving the
probabilistic expression of behaviors among individuals.
For example, an individual under a mixed strategy could
express one mating tactic, such as sneaking, with a certain frequency and another tactic, such as mate guarding,
at all other times.[3] Though a mixed strategy is theoretically possible, it has not been documented in the context
of alternative mating behaviors. Instead, a conditional
strategy involving alternative behaviors may best characterize alternative mating strategies.[1]
Condition-dependent behavior in the context of mating
may result from changes in resource availability and intrasexual competition for mates. When competition decreases, the expression of alternative behaviors also decreases. Changes in mating behaviors, especially among
alternative males, have been documented in insects, sh,
and amphibians upon removal of dominant males.[3] Additionally, the availability of mates and resources also af-

2.2

Female mimicry by males

fects the expression of alternative strategies within a sex.

The gain or loss of territory has been shown to aect mating approaches among insect species, while the receptivity and spatial distribution of mates impacts tactics used
among insects, sh, and mammals.[3] Mating behaviors
are also aected by an individuals size and age, as smaller
or younger individuals are more likely to attempt reproduction through alternative means, including mimicry or
sneak tactics.[3] As a result, the ability to choose a behavior that maximizes tness under certain circumstances
evolves.[1]

3
for the males practicing them, and adoption of these alternative mating strategies has contributed to the maintenance of a dimorphic male population.[2]
2.1.2 High-backed pygmy swordtail (Xiphophorus
multilineatus)

Pygmy swordtail males oer another example of alternative mating strategies. Some males mature later at a larger
size and always use courtship behavior, while other males
mature early at a smaller size, sometimes using courtship
behavior when alone with a female, but more often using
sneaky behavior. This behavior is not preferred by the
2 Alternative male strategies
female, and is therefore not as successful as courtship in
gaining matings, however the higher probability of surIt has long been known that males in a wide variety of viving to reach sexual maturity due to maturing early is
the smaller sneaker males in the
animal populations practice alternative mating strategies enough to maintained
[5]
population.
in order to maximize their reproductive tness. This is
especially common when there is male-male competition
for access to mates. In cases where such alternative strate- 2.1.3 Red paper wasps (Polistes canadensis)
gies are as successful at obtaining mates as the predominant strategy, a coexistence of dierent mating strategies Male red paper wasps engage in the role of the patroller
will evolve. Below are a few common examples of male as an alternative mating tactic to the role of the territoalternative mating strategies.
rial male (who chases away intruders). Patrollers have a

2.1

Sneaking behavior in males

Sneaking behavior may refer to a strategy that allows a

male to more stealthily access a female partner, often
avoiding altercations with other more dominant males.

2.1.1

Horned beetles (Onthophagus acuminatus)

Horned beetles demonstrate alternative mating strategies

due to dierent nutritious conditions during development
that aect adult body size. In this species, males who
receive high levels of nutrition during development will
surpass a size threshold above which they develop large
horns. Males who do not pass the threshold will develop
either small or nonexistent horns. These varying phenotypes will lead individual males to adopt dierent mating
strategies. Those who develop long horns will practice
mate guarding, protecting the entrance to the tunnel in
which a female is resting or feeding. These males will
ght any male that attempts to enter. This is a common
strategy observed in populations in which females are dispersed and have synchronized periods of fertility, as well
as those in which females are found in clusters that can
be guarded to maintain access to more than one female.
Smaller males with little or no horns have little chance of
beating larger males in altercations and will thus adopt an
alternative sneaking strategy, digging a new tunnel that
will allow them to intercept the females tunnel without
being noticed by the guarding male. Both of these strategies have proven, thus far, to be reproductively eective

smaller body size than territorial males. There is significant competition over the possession of territories. Although these territories do not necessarily hold any resources or nesting sites, owning a territory tends to lead to
a greater number of mating opportunities. Males attract
females to these territories by rubbing their abdomens
across the territories to apply pheromones. Because of
their inability to successfully compete against the larger
territorial males for territories, smaller males resort to patrolling. But patrollers do not just wait around for territories to be vacated; they will sneak matings with females
in territories when the territorial males are temporarily
away or distracted. [6]

2.2 Female mimicry by males

Males practicing female mimicry may do so in order to
gain access to mates in areas of where only females congregate.
2.2.1 Marine isopod
In Paracerceis sculpta there are three genetically distinct
male morphs. Alpha males, which represent the largest
and most common male morph, tend to defend harems in
order to monopolize access to a large number of females.
This is the predominant mating strategy in this species.
Beta males are about the same size as female isopods, and
they take advantage of that fact by mimicking female behavior in order to enter harems and gain access to fertile
females. Gamma males are the smallest morph. These individuals adopt a sneaking strategy and rely on their small

4 SEE ALSO
circumvent these limitations. Below are some examples
of alternative female strategies seen in nature.

3.1 Copying mate choice

In the guppy, Poecilia reticulata, females will copy another females mate choice if given the opportunity to
watch the other female choose. While older females
do not copy younger females, younger females will copy
older females.[10][11] This copying behavior arises from a
dierence in ability to assess males. Since this behavior
only arises when in the presence of another female, it is a
behavioral alternative to the norm of just choosing a male
mate based on personal assessment.

3.2 Sneaking behavior in females

Paracerceis sculpta illustration

body size to enter harems undetected and remain in them

while they seek mating opportunities.
These very distinct strategies, all determined by a single
genetic locus, give equivalent lifetime mating success to
each of the three morphs, indicating that natural selection is not acting on one morph more strongly than another. All three alleles expressed in the marine isopod
population will continue to contribute to male morphology as long as the reproductive success granted by each
one continues to be as benecial as the others.[7]

In the damselsh, Chromis multilineata, females can often become infected with the parasite Anilocra chromis.
In the event of infection, males do not allow infected females into the nest and do not mate with them. Thus, to
bypass this limitation to mating, infected females will often sneak into male nests.[12] Although the female is often
immediately chased out, this behavior serves as evidence
that sneaking is not just an alternative male strategy. In
fact, sneaking is just a common strategy for any sex that is
denied mating to a certain class of animals. The strategy
of these infected females is therefore another behavioral
alternative strategy.

3.3 Male mimicry by females

Alternative female strategies

Historically, while male alternative strategies have been

well documented, alternative female strategies have not
been studied extensively. This large discrepancy in information is mostly due to two factors. First, male mating behavior is typically driven by competition for mates,
such as physical competition, territoriality, or parental
care investment. Thus, male alternative behaviors arise
as a direct result of these various forms of competition. However, females typically do not compete directly for these resources or mates. Instead, females indirectly compete through dierences in premating, mating
and post-mating behavior.[8] The subtle nature of female
competition makes alternative behaviors very dicult to
study relative to males. Second, males are more likely
to experience sexual selection than females. Due to this
increased selection, it is statistically more likely for alternative strategies to evolve in males than females.[9] However, though subtle and slightly more rare, females are still
major participants in the mating process and can also experience limitations in access to males and male parental
care. Thus, alternative female strategies have evolved to

In damselies, Ischnura, females are frequently harassed

by males that wish to mate. There is signicant variation in the females physical abilities to tolerate male
mating harassment. In this species, there is a physical
dimorphism: one type is cryptic (heteromorphic) and the
other type looks like a male (andromorph). In many cases
the andromorph even behaves like a male when among
other males. Studies have found that the andromorph
only mates half as often as the heteromorph.[13] While
a decrease in mating would be devastating for males, it
is often an advantage in females. For females, excessive
mating is a waste of time and energy and it increases exposure to predators. Thus, the ability to ward o extra
mating gives the andromorphs a frequency dependent selective advantage. This is example of a traditionally male
characterized Mendelian alternative strategy that has now
been observed in females.

4 See also
Mating system

References

[1] Gross, Mart R. (1996). Alternative reproductive strategies and tactics: diversity within sexes. Tree 11 (2):
9298. doi:10.1016/0169-5347(96)81050-0. PMID
21237769.
[2] Pagel, Mark D. Mating Strategies, Alternative. Encyclopedia of Evolution. Oxford UP.
[3] Dominey, Wallace J. (1984). Alternative mating tactics
and evolutionarily stable strategies. American Zoology
24: 385396. doi:10.1093/icb/24.2.385.
[4] Shuster, Stephen M.; Michael J. Wade (2003). Mating
Systems and Strategies. Princeton University Press. pp.
434450. ISBN 9780691049311.
[5] Molly R. Morris, Oscar Rios-Cardenas, Jason Brewer.
Variation in mating preference within a wild population
inuences the mating success of alternative mating strategies, Animal Behaviour, Volume 79, Issue 3, March 2010,
Pages 673-678
[6] Polak, Michal. Competition for Landmark Territories among Male Polistes canadensis (L.) (Hymenoptera:
Vespidae): Large-size Advantage and Alternative Mateacquisition Tactics. Behavioral Ecology 4.4 (1993): 32531.
[7] Equal Mating Success among Male Reproductive Strategies in a Marine Isopod. Shuster, Stephen M; Wade,
Michael J. Nature; Apr 18, 1991; 350, 6319; ProQuest
Research Library pg. 608
[8] Ne, BD; Svensson EI (2013). Polyandry and alternative
mating tactics. Phil Trans R Soc (B368).
[9] Henson, SA; Warner RR (1997). Male and female alternative reproductive behaviors in shes: a new approach
using intersexual dynamics. Annu Rev Ecol Syst 28: 571
92. doi:10.1146/annurev.ecolsys.28.1.571.
[10] Dugatkin, LA; Godin J-GJ (1992). Reversal of female mate choice by copying in the guppy Poecilia reticulata". Proc R Soc London Ser B 249 (1325): 17984.
doi:10.1098/rspb.1992.0101.
[11] Dugatkin, LA (1992). Sexual selection and imitation: females copy the mate choice of others. Am Nat 139 (6):
138489. doi:10.1086/285392.
[12] Johnston, BA (1996). The pathological and ecological consequences of parasitism by a cymothoid isopod
(Anilocra chromis) for its damselsh host (Chromis multilineata)". MS thesis. Univ Calif Santa Barbara: 81.
[13] Fincke, OM (2004). Polymorphic signals of harassed female odonates and the males that learn them support a
novel frequency-dependent model. Animal Behavior 67
(5): 83345. doi:10.1016/j.anbehav.2003.04.017.

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